Molecular Systematics of New World Suboscine Birds R
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Available online at www.sciencedirect.com MOLECULAR SCIENCEENCE^I /W) DIRECT® PHYLOGENETICS AND EVOLUTION ELSEVIER Molecular Phylogenetics and Evolution 32 (2004) 11-24 www.elsevier.com/locate/ympev Molecular systematics of New World suboscine birds R. Terry Chesser* Department of Ornithology, American Museum of Natural History, Central Park West at 79 th Street, New York, NY 10024, USA Received 12 February 2003; revised 14 November 2003 Available online 4 February 2004 Abstract Pliylogenetic relationships among New World suboscine birds were studied using nuclear and mitochondria! DNA sequences. New World suboscines were shown to constitute two distinct lineages, one apparently consisting of the single species Sapayoa aenigma, the other made up of the remaining 1000+ species of New World suboscines. With the exception oí Sapayoa, monophyly of New World suboscines was strongly corroborated, and monophyly within New World suboscines of a tyrannoid clade and a furnarioid clade was likewise strongly supported. Relationships among families and subfamilies within these clades, however, differed in several respects from current classifications of suboscines. Noteworthy results included: (1) monophyly of the tyrant- flycatchers (traditional family Tyrannidae), but only if the tityrines (see below) are excluded; (2) monophyly of the pipromorphine flycatchers (Pipromorphinae of Sibley and Ahlquist, 1990) as one of two primary divisions of a monophyletic restricted Tyrannidae; (3) monophyly of the tityrines, consisting of the genus Tityra plus all sampled species of the Schiffornis group (Prum and Lanyon, 1989), as sister group to the manakins (traditional family Pipridae); (4) paraphyly of the ovenbirds (traditional family Furnariidae), if woodcreepers (traditional family Dendrocolaptidae) are excluded; and (5) polyphyly of the antbirds (traditional family Formi- cariidae) and paraphyly of the ground antbirds (Formicariidae sensu stricto). Genus Melanopareia (the crescent-chests), although clearly furnarioid, was found to be distant from other furnarioids and of uncertain aflSnities within the Furnarii. Likewise, the species Oxyruncus cristatus (the Sharpbill), although clearly tyrannoid, was distantly related to other tyrannoids and of uncertain aflSnities within the Tyranni. Results of this study provide support for some of the more novel features of the suboscine phylogeny of Sibley and Ahlquist (1985, 1990), but also reveal key differences, especially regarding relationships among suboscine families and subfamilies. The results of this study have potentially important imphcations for the reconstruction of character evolution in the suboscines, especially because the behavioral evolution of many suboscine groups (e.g., Furnariidae) is of great interest. © 2003 Elsevier Inc. All rights reserved. 1. Introduction sively in the tropics of the Old World, where they appear to be relictually distributed (Mayr and Amadon, 1951). Suboscine, or non-oscine, birds are one of two major Although suboscines were defined in part based on components of tlie large avian order Passeriformes lack of a character•they are passerine birds that lack (passerines) and consist of roughly 1150 species, or the oscine, or songbird, syrinx•and have been referred about one-eighth of extant birds. Found throughout the to as "this by no means natural group" (Stresemann, Americas, they account for more than 30% of the 1934, translated in Sibley, 1970), the modern view is that world's richest avifauna, that of the Neotropics, where suboscines are monophyletic relative to other passerines they have undergone a remarkable large-scale radiation. and that they consist of two monophyletic groups Suboscines are predominantly a New World group, but divisible along geographical lines: the New World su- the 51 species of the families Eurylaimidae (broadbills), boscines and the Old World suboscines (Irestedt et al., Philepittidae (asities), and Pittidae (pittas) occur exclu- 2001; Raikow, 1987; Raikow and Bledsoe, 2000; Sibley and Ahlquist, 1990). New World suboscines are likewise generally con- Present address: Australian National Wildlife Collection, CSIRO Sustainable Ecosystems, GPO Box 284, Canberra ACT 2601, Austra- sidered to consist of two monophyletic groups, one lia. Fax: 1-61-2-6242-1688. ("Tyranni" in Ames, 1971 and Raikow, 1987, "Tyr- E-mail address: [email protected]. annida" in Sibley and Ahlquist, 1990) consisting of the 1055-7903/$ - see front matter © 2003 Elsevier Inc. All rights reserved. doi:10.1016/j.ympev.2003.11.015 12 R T. Chesser I Molecular Phylogenetics and Evolution 32 (2004) 11-24 tyrant-flycatchers and relatives, the other ("Furnarii" of New World suboscines, and (3) the relationships of in Ames, 1971 and Raikow, 1987, "Furnariida" and suboscine taxa of uncertain affinities. "Thamnophilida" in Sibley and Ahlquist, 1990) con- sisting of the ovenbirds, antbirds, and related taxa. The Tyranni contain many taxa of uncertain familial affini- 2. Materials and methods ties, variously placed in the traditional families Tyran- nidae (tyrant-flycatchers), Pipridae (manakins), or Fifty-three taxa were sampled (Table 1), including Cotingidae (cotingas), including members of the Schif- one non-passerine (the woodpecker Campethera nivosa), fornis group (Prum and Lanyon, 1989) and the Sharpbill four oscine passerines (two representatives of each os- Oxyruncus cristatus. Species composition of families in cine parvorder in Sibley and Ahlquist, 1990), one pas- the Furnarii (sensu Ames, 1971 and Raikow, 1987), in serine of uncertain affinities (the New Zealand wren contrast, has with few exceptions been well defined, but Acanthisitta chloris), three Old World suboscines (one the monophyly and relationships of specific families representative from each family), and 44 New World have been a matter of debate. suboscines. Taxon sampling among New World subos- Although most previous systematic work on subos- cines was guided largely by the phylogeny of Sibley and cines has been based on morphology, Sibley and Ahl- Ahlquist (1985, 1990) and the classification of Sibley and quist (1985, 1990) produced a detailed phylogenetic Monroe (1990), and was designed to establish relation- hypothesis of New World suboscines based on DNA- ships among major groups, to provide simple tests of the DNA hybridization data (Fig. 1). This phylogeny was monophyly of major groups, and to indicate relation- consistent in some ways with traditional classifications, ships of problematical taxa. The following were con- but it contained several striking features, including sidered major groups for sampling purposes, using the polyphyly of the traditional family Tyrannidae (species nomenclature of Sibley and Monroe (1990), with num- included in the Tyranninae and Pipromorphinae of ber of taxa sampled in parentheses: Pipromorphinae (5), Sibley and Ahlquist) and polyphyly of the traditional Tyranninae (4), Tityrinae (3), Cotinginae (5), Piprinae family Formicariidae (species included in the Formi- (3), Furnariinae (6), Dendrocolaptinae (3), Thamno- cariidae and Thamnophilidae of Sibley and Ahlquist). philidae (4), Formicariidae (4), Conopophagidae (2), Polyphyly of the traditional Formicariidae has also been and Rhinocryptidae (4). The Broad-billed Sapayoa, supported by a recent study using DNA sequence data Sapayoa aenigma, a New World suboscine of uncertain (Irestedt et al., 2002). affinities, was also sampled. Larger numbers of taxa Below I provide a phylogenetic hypothesis for New were sampled for groups with relatively large numbers World suboscines based on DNA sequence data, and of species (e.g., Furnariidae) or for more controversial use this hypothesis to address: (1) the monophyly of groups (e.g., Pipromorphinae), and smaller numbers for New World suboscines, (2) the monophyly and relatively small groups (e.g., Conopophagidae). Taxa relationships of traditional and non-traditional groups within major groups were chosen to be as divergent as possible, to strengthen tests of monophyly and to break OW suboscines up long branches for phylogenetic analysis. Intron 7 of the nuclear gene ß-fibrinogen and the Tyranninae complete mitochondrial genes ND3 and COII were se- Tityrinae quenced for all taxa, with two exceptions. Complete mitochondrial but only partial ß-fibrinogen sequence Cotinginae was obtained for Melanopareia torquata, and complete ß-fibrinogen but no mitochondrial sequence was ob- Piprinae tained for A. chloris. Tissue samples were collected Pipromorpliinae during fieldwork in South America and from the De- partment of Ornithology, American Museum of Natural Furnariidae History, New York, NY; the Genetic Resources Col- lection, Louisiana State University Museum of Natural Formicariidae Science, Baton Rouge, LA; the Division of Birds, Field Museum of Natural History, Chicago, IL; the Royal Rliinocryptidae Ontario Museum, Toronto, Ont.; the Marjorie Barrick Conopophagidae Museum, University of Nevada Las Vegas, Las Vegas, NV; and the University of Arizona Bird Collection, Thamnoplniiidae Tucson, AZ. Sequences were obtained using DNA ex- Fig. 1. Phylogenetic hypothesis of suboscine birds based on DNA- tracted from tissue by means of a 5% Chelex solution DNA hybridization data (Sibley and Ahlquist, 1985, 1990). This is (Walsh et al., 1991). Primers used for initial PCR Sibley and Ahlquist's preferred tree, obtained using UPGMA analysis. amplification of intron 7 were FIBI7U and FIBI7L, both R. T. Chesser I Molecular Phylogenetics and Evolution 32 (2004) 11-24 13 Table 1 List